The target article misunderstands the research program it criticizes. The work of Boyd, Richerson, Fehr, Gintis, Bowles and their collaborators has long included the theoretical and empirical study of models both with and without diffuse costly punishment. In triaging the situation, we aim to (1) clarify the theoretical landscape, (2) highlight key points of agreement, and (3) suggest a more productive line of debate.

Over the past several decades, we have argued that cultural evolution can facilitate the evolution of large-scale cooperation because it often leads to more rapid adaptation than genetic evolution, and, when multiple stable equilibria exist, rapid adaptation leads to variation among groups. Recently, Lehmann, Feldman, and colleagues have published several papers questioning this argument. They analyze models showing that cultural evolution can actually reduce the range of conditions under which cooperation can evolve and interpret these models as indicating that we were wrong to conclude that culture facilitated the evolution of human cooperation. In the main, their models assume that rates of cultural adaption are not strong enough compared to migration to maintain persistent variation among groups when payoffs create multiple stable equilibria. We show that Lehmann et al. reach different conclusions because they have made different assumptions. We argue that the assumptions that underlie our models are more consistent with the empirical data on large-scale cultural variation in humans than those of Lehmann et al., and thus, our models provide a more plausible account of the cultural evolution of human cooperation in large groups.

The use of socially learned information (culture) is central to human adaptations. We investigate the hypothesis that the process of cultural evolution has played an active, leading role in the evolution of genes. Culture normally evolves more rapidly than genes, creating novel environments that expose genes to new selective pressures. Many human genes that have been shown to be under recent or current selection are changing as a result of new environments created by cultural innovations. Some changed in response to the development of agricultural subsistence systems in the Early and Middle Holocene. Alleles coding for adaptations to diets rich in plant starch (e.g., amylase copy number) and to epidemic diseases evolved as human populations expanded (e.g., sickle cell and deficiency alleles that provide protection against malaria). Large-scale scans using patterns of linkage disequilibrium to detect recent selection suggest that many more genes evolved in response to agriculture. Genetic change in response to the novel social environment of contemporary modern societies is also likely to be occurring. The functional effects of most of the alleles under selection during the last 10,000 years are currently unknown. Also unknown is the role of paleoenvironmental change in regulating the tempo of hominin evolution. Although the full extent of culture-driven gene-culture coevolution is thus far unknown for the deeper history of the human lineage, theory and some evidence suggest that such effects were profound. Genomic methods promise to have a major impact on our understanding of gene-culture coevolution over the span of hominin evolutionary history.

Much of human cooperation remains an evolutionary riddle. Unlike other animals, people frequently cooperate with non-relatives in large groups. Evolutionary models of large-scale cooperation require not just incentives for cooperation, but also a credible disincentive for free riding. Various theoretical solutions have been proposed and experimentally explored, including reputation monitoring and diffuse punishment. Here, we empirically examine an alternative theoretical proposal: responsibility for punishment can be borne by one specific individual. This experiment shows that allowing a single individual to punish increases cooperation to the same level as allowing each group member to punish and results in greater group profits. These results suggest a potential key function of leadership in human groups and provides further evidence supporting that humans will readily and knowingly behave altruistically.

If individuals will cooperate with cooperators, and punish non-cooperators even at a cost to themselves, then this strong reciprocity could minimize the cheating that undermines cooperation. Based upon numerous economic experiments, some have proposed that human cooperation is explained by strong reciprocity and norm enforcement. Second-party punishment is when you punish someone who defected on you; third-party punishment is when you punish someone who defected on someone else. Third-party punishment is an effective way to enforce the norms of strong reciprocity and promote cooperation. Here we present new results that expand on a previous report from a large cross-cultural project. This project has already shown that there is considerable cross-cultural variation in punishment and cooperation. Here we test the hypothesis that population size (and complexity) predicts the level of third-party punishment. Our results show that people in larger, more complex societies engage in significantly more third-party punishment than people in small-scale societies.

Human morality is a key evolutionary adaptation on which human social behavior has been based since the Pleistocene era. Ethical behavior is constitutive of human nature, we argue, and human morality is as important an adaptation as human cognition and speech. Ethical behavior, we assert, need not be a means toward personal gain. Because of our nature as moral beings, humans take pleasure in acting ethically and are pained when acting unethically. From an evolutionary viewpoint, we argue that ethical behavior was fitness-enhancing in the years marking the emergence of Homo sapiens because human groups with many altruists fared better than groups of selfish individuals, and the fitness losses sustained by altruists were more than compensated by the superior performance of the groups in which they congregated.

Why Humans Cooperate takes a unique look at the evolution of human cooperation and tries to answer the question: why are people willing to help others at a cost to themselves? The book brings together evolutionary theories, economic experiments, and an anthropological case study that runs throughout the book to explain and illustrate human cooperation.

Using an evolutionary framework, Natalie and Joseph Henrich have expanded upon several diverse theories for explaining cooperative or ‘helpful’ behavior, and integrated them into a unified theory. Established concepts such as kin selection and reciprocity have been linked with theories on social learning and our evolved psychologies to explain the universality of human cooperation—as well as the distinctive ways in which cooperative behavior expresses itself in different cultures.

The theories developed in the book are brought to life by examining the Chaldeans of metropolitan Detroit. By exploring Chaldean cooperation, theoretical concepts are shown to translate into social behavior, and universal psychologies for cooperation lead to culturally-specific norms, beliefs, and practices. The book also introduces a series of economic experiments that help us understand why, when, and to what extent people are willing to help others. These experiments also highlight the variation in behaviors across cultural groups, even when all the groups rely on the same cognitive machinery and evolved psychologies. The merging of theory, experiments, and the Chaldean case study allows for an in-depth exploration of the origins and manifestations of cooperation.